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Home , Black caracara, Cathartes, Crested caracara, King vulture, Mauritia flexuosa

458 SHORT COMMUNICATIONS VOL. 39, NO. 4 j. RaptorRes. 39(4):458-461 ¸ 2005 The Raptor ResearchFoundation, Inc.

KING (SARCORAMPHUS PAPA) FORAGE IN MORICHE AND CUCURIT PALM STANDS

MARSHA A. SCHLEE1 MusgumNational dZIistoire Naturelie, Dgpartement t•cologie etGestion dela Biodiversitg,USM 0305, CP 31 Mgnagene, 57 rue Cuvig 75231 Paris cedex05, France

KEYWORDS: Kit'•gVulture,, Sarcoramphus papa; batana). The moriche palms ()can be flexuosa;Attalea maripa palms;wedge-capped capuchin mon- found scatteredin the gallery forests or in stands (mor- keys;Cebus olivaceus;foraging association. ichales;see GonzftlezBoscfm 1987) in the seasonallyIn- undated areasof the valley.The morichefruits, 3-7 cm long, ovate to globular and having an oily mesocarp Feeding on palm fruit, particularly drupes of the Af- (Borgtoft Pedersenand Balslev1990), fall to the ground rican oil palm (Elaeisguineensis), has been documented when almost ripe and accumulate in the water among for severalOld World speciesof of prey (Thiollay fallen fronds and debris. Cucurit palms (Attaleamaripa = 1978, Barlow 2004). In the New World, fruits of the im- Maximilianaregia) occur as standswithin the gallery for- ported African oil palm havebeen consumedby the Tur- estson dry terrain. The fruits, ovate,5-7 cm long, are key ( ruficollis;Pinto 1965), Yellow- alsorich in oil (Braun 1997). The observationsreported headed (Milvago chimachima;Haverschmidt here took place during the rainy season,which lastsApril 1962), and BlackVulture (C0ragypsatratug Haverschmidt through October or November. The study periods (24 1947, Pinto 1965, Elias and Valencia 1982). Both vulture June-24 July 1994, 26 June-14 August 1995, 28 May-14 July 1996, 30 June-18 August 1997) were set up to co- species,as well as the (Caracara cheri- incide with the fruiting seasonreported locallyfor both way), consume flesh of coconuts (Cocosnucifera; Hav- palm species.Observations were carried out daily, gen- erschmidt1947, Crafts 1968), and fruits of palms (Maur- erally from 0615-1900 H. The length of each monitoring ztzaflexuosa and Desmoncussp.) have been found in period (up to 5 hr) depended on weather,logistics, and stomach contents of the Black Caracara (Daptrius at• whether or not the King Vultures were present in the Haverschmidt 1962). American Black Vultures also feed valley.The number of sample daysfor each studyperiod on sweetpotatoes (McIlhenny 1945) and avocado(R6hl varied from 29-50 d, with the afternoon of arrival and 1949) when is scarce,and Turkey Vultures ingest morning of departurebeing countedif observationswere leaves, seeds, and bark of cottonwood trees (Populus carried out (Table 1). I used 10X binoculars and ob- spp.), apparently as casting material (Davis 1983). No servedfrom outcropsand other high points.

published data were found on ingestionof matter RESULTS by King Vultures (Sarcoramphuspapa), but residentsat Hato Las Nieves, reported that the species In all four years,moriche and cucurit fruit production consumesfruits of the moriche (Mauritia flexuosa)and took place earlier than expected and little remained by cucurit (Attaleamaripa) palms when carrion is scarce(Y. mid-July.Both palm speciessuffered from drought in Carbonell and A. Mendoza pers. comm.). The observa- 1995 and 1996. A mean of 2.1 -+ 1.1 (SD) King Vultures tions reported in this paper were gathered in an attempt (range = 1-4) foraged in four different morichales (N to verify these claims. = 7 occurrences),and 3.4 _ 1.0 (range = 2-4) foraged (N = 7 occurrences)in two of the cucurit stands,pri- STUDY AREA AND METHODS marily during the 1994 and 1995 field seasons(Table 1). My observationswere part of a long-term study (1994- This activitywas most often carried out by two adults to- 2000) on the abundance, population structure, move- gether (N -- 5) or three adults and an immature (N = ment patterns, and foraging strategiesof King Vultures m the Serrania de la Cerbatana, Estado Bolivar, Venezue- 6), presumed to be the local birds. Eleven of the 14 oc- la The studywas conductedin the southeasternpart of currencestook place when the vultures had not fed on the massifat Hato Las Nieves (SabanaNueva: 6ø34'80"N, livestock carcasses(natural mortality, including 66ø12'17"W), a ranch located ca. 125 km south of Caicara [Pantheraonca] predation) or inedible parts of slaugh- del Orinoco. The valley of Las Nieves, ca. 20 km long tered for 2-3 wk. Four bouts in the cucurit stands (northwest-southeast)and 9 km wide, is dominated by (August 1995) followed presumed feeding on the re- lowland shrub ,mainly 220-260 m abovesea lev- mains of jaguar-killed native wildlife. el. The bordering mountainsreach elevationsof 1600 m On 27June 1994, at 0947 H, four AmericanBlack Vul- to the west and 1880 m to the north (Cerro de la Cer- tures flew from the western morichal. As I approached, I sightedan adult perched low at the edge • Email address: [email protected] of the palm standand another adult on the ground near- DECEMBER 2005 SHORT COMMUNICATIONS 459

Table 1. Mean number of King Vultures per studyperiod (1994-97) seen foraging in moriche and cucurit palm standsand number of occurrencesat Hato Las Nieves, Venezuela. Number of days the vultures fed on livestock carcassesand total daysof observationare given.

NUMBER OF DAYS NUMBER OF FREQUENCY STUDY KING VULTURES LIVESTOCK PEmOD • +-- SD (RANGE) MOmCHE CUCUmT CARCASSES OBSERVATION

1994 2.2 +-- 1.0 (1--4) 3 3 15 29 1995 3.3 --+ 1.3 (1--4) 3 4 11 50 1996 3.0 + 0 (3) 1 0 26 48 1997 0 0 0 15 49

by at the base of the only fruiting moriche in the area. ed, I found severalfruits with the mesocarpwholly or This , with head down, just out of sight, seemed to partially scrapedoff. No carrion was present, but a me- be nibbling on something.When both birds flew to near- dium-sized savanna tortoise (Geochelonecarbonaria) was in by palms, I could see that their crops were extended. the area and could have been feeding on the fruits ear- Their flight attracted a that walked lien Later (1157 H), I locatedthe immaturebird feeding around the area but did not feed. At the site, I found at the base of another fruiting moriche. Again, the me- one unripe moriche fruit with half the mesocarpfreshly socarpon severalfruits had been freshly scrapedoff. No scrapedoff longitudinallyon one side, the marks clearly debris or carrion was found, and no live animals were imprinted on the nut, and a large piece of overripe me- nearby.I concluded that the King Vulture had eaten the socarpfrom another fruit. No debris, no carrion, and no pulp. Two dayslater, at 0740 H, an adult perched in the live animals were present. I concluded that the King Vul- gallery forest north of the central camp, and at 0913 H ture had consumedthe missingpulp, but this would not I found it foraging on the ground among the few palms explain crop extension. The next day after the rains that remained of a remnant morichal. No carrion was stopped (1620 H), I observed an adult sunning in the present. On 29 July 1995, 8 d after a carcasshad been main cucurit stand.As I penetrated into the gallery for- consumed, two adults suddenly flew up from one of the est, I came acrossanother King Vulture rummaging in gallery-forestfloors (1645 H) in an area that had fruiting the litter at the base of a cucurit palm. The debris con- moriche, but I was unable to investigatefurther. Then, rained no matter, only cucurit fruits, whole or rot- on 5-6 and 9-10 August 1995, three adults and an im- ting or partially eaten, piecesof mesocarp,and clean ker- mature, presumed to be the same birds as above, were nels. Several wedged-capped capuchin monkeys (Cebus sighted in a cucurit stand. Eight Turkey Vultures were ohvaceus)were in the stand. Two days later (30 June also present. The King Vultures perched within the up- 1994), two adults were observed in a morichal at ca. 1100 per strata of the canopy of the broad-leafedtrees, occa- H and in a cucurit stand in the afternoon, and the fol- sionallygoing to the top of the trees to sun or dry, and lowing day three adults and an immature were in the spent the mornings in the stand.A troop of ca. 30 wedge- same cucurit stand at mid-day. Capuchin monkeyswere capped capuchin monkeys,with many femalestransport- present both days.The last sightingin 1994 took place ing infants, was foraging there on the first two days.The on 7 July, 2 d after the vultures had consumeda dead area was strewn with large pieces of bark and fallen horse.At 0737 H an adult was again perched low at the branches. On the last 2 d, fewer capuchinswere present, edge of a morichal. It was raining hard (1000 H) when but were seen with a pair of red-howler monkeys (Al- I found the adult foraging under the only fruiting mor- ouatta seniculus). I found no remains of carrion. lche in the area. The bird flushed upon seeing me, but On 8June 1996 at 0913 H, 4 d after the vultures had then returned to the morichal again, weavingin and out eaten livestockcarrion, I came upon two adult King Vul- of the vegetationand foraging on the ground asI tracked tures and an immature resting on a low shrub at the edge it for ca. 90 min. Moriche fruits were present in the areas of a morichal and next to a fruiting moriche palm. Three where the bird had foraged. I found no carrion or live American Black Vultures were feeding on the ground at animals. The vulture's crop was extended, indicating it the base of the palm. The crop of one King Vulture was had ingestedfood while in the morichal. slightlyextended. I found no carrion and no live animals, On 28 June 1995, at 0722 H, two adult King Vultures only palm fruits lacking part of the mesocarpand show- and an American perched at the edge of ing signsof having been scraped. I concluded that both a morichal. An hour later another adult King Vulture and vulture specieshad been feeding on the fruits. This was an immature joined them, and one of the first adultsflew the only time the King Vultures were observedto forage to the ground near a palm with fruit. When I investigat- in a morichal in 1996, but very few moriche palms were 460 SHORT COMMUNICATIONS VOL. 39, NO. 4 fruiting (-<10% in the major stands).No foraging took the movementsof monkey troops (e.g., arboreal : place in the cucurit stands,but only two cucurit palms Zhang and Wang 2000). had fruit, althoughfermenting drupeswere found at the How could King Vulturesbenefit from associatingwith base of others. Livestock carrion was abundant (Table 1), Cebusmonkeys? Is it only to profit from occasionalpri- and were in the area. mate mortality?Although 55% of the wedge-cappedca- The King Vultureswere not seenforaging in the palm puchin's diet consistsof plant matter, particularly ripe standsin 1997, even though more moriche and cucurit fruits, invertebratesare searchedout by peeling off loose were fruiting than in 1996. A small troop of capuchin bark, digging into rotting material and sifting through monkeyswas present twice in the studyarea, and domes- leaf debris on the ground (Robinson 1986). Perhapsthe tic carrion was not often available (Table 1), as most live- King Vulturesbenefit from larvaethat are exposedor fall stock had been removed from the valley. The vultures to the ground while the troop forages; vultures are pmkedover the debrisat former carcasssites on 12 d and known to scoopup maggotsfrom decomposingcarcasses followed a jaguar. This feline was known to have come (Houston 1988). Of greater interest is the occasional through the valleyon four occasions,and the King Vul- predatorybehavior of the capuchinson vertebrates.For tures presumablyfed on the remains of kills on native example, the visceraof lizardsmay be eaten but the mus- wfidlife at or near Las Nieves on 5 d. cular part left, and the remains of captured frogs d•s- carded (Robinson 1986). In Cebuscapucinus, after wres- DISCUSSION tling with an Iguanasp., a monkeymanaged to break off 30-40 cm of the iguana'stail, stripped some meat from My observationssupport the claimsthat the King Vul- it, and droppedthe rest (Baldwinand Baldwin1977). By tures at Hato Las Nieves eat fruits of the moriche palm, followingwedge-capped capuchin monkeys, the vultures particularlywhen carrion is scarce.Eating moriche fruits could glean the remains of discarded vertebrate prey. may partially compensatethe lack of carrion, 100 g of However, eating moriche fruit or monitoring monkeys fresh pulp having10.5 g of fat and 3.0 g of protein, but did not seem to be as beneficial as picking over the scat- the fruits could havebeen ingestedfor their vitamin-min- tered remains of former carcassesand followingjaguars eral content (see Gonzalez Bosc•m1987, Borgtoft Ped- to consume the remains of kills. These observations fi- ersen and Balslev 1990). From the remains of the fruits lustrate some opportunistic feeding strategiesused by found at the feeding sites,I conclude that the mesocarp King Vultures. was scrapedoff longitudinally,but judging from the size of extendedcrops of somevultures, some fruits, probably SARCORAMPHUSPAPA FORR•/EA EN MORICHALESYEN GRUPOS the smaller ones, may have been swallowedwhole. Both DE PALMAS DE CUCURTO feedingtechniques are usedby the Palm-nutVulture (Gy- pohieraxangolensis) when consumingdrupes of the Afri- RESUMEN.--Estetrabajo se realiz6 durante las 6pocas11u- can oil and Raphiapalms, the ingestedkernels being re- viosasdesde finales de junio de 1994 hasta mediados de gurgitated later (Thiollay 1978). Only once in my agostode 1997 en la Serraniade la Cerbatana,Hato Las observationswas a potential fallen-fruit consumer pre- Nieves, Estado Bolivar, Venezuela. Los datos recolectados sent--a tortoise. My observationsalso lend support to sostienenlas afirmacionesdel personaldel hato de que claims that American Black Vultures eat Mauritia fruits Sarcoramphuspapas come frutos de la palma moriche (Y. Carbonell and A. Mendoza pers. comm.). Moriche (Mauritia fiexuosa),principalmente cuando escaseala ca- rrofia. Una media de 2.1 individuosde S. papa (rango = fruit-eating appearsto be an activitycarried out by vul- 1-4) forrajearon en los morichales(N = 7 avistamien- tures local to the study area; however,because north- western Bolivar state is one of the t•w areas in the Ve- tos). No pude confirmar las alirmacionesde que S. papa come tambi6n fi:utos de la palma de cucurito (Attalea nezuelan Guayana that has a high concentration of maripa).Los fi:utosde ambasespecies de palma contie- morichales (see Gonzalez Boscfin 1987), moriche fruit- hen aceite. En mis obscrvaciones, una media de 3.4 m- eating by King Vultures could be more widespreadthan at Hato Las Nieves. dividuos (rango = 2-4) se encontraronvigilando tropas del mono capuchinoCebus olivaceus que habian venido a On the other hand, I wasnot able to confirm that King forrajear a los rodalesde cucuritos.Sugiero que S. papa Vultures eat cucnrit fruits, although their consumptionis podria asociarsecon los monos para aprovecharlos in- plausible consideringthe oil content of the mesocarp vertebradosque espantan,como las larvas de insectos, (Braun 1997). Whenever the King Vultures lbraged in los restosde vertebradosque capturany ocasionalmente the cucurit stands, wedge-capped capuchin monkeys los cad/tveres de toonos. were also present,which suggeststhat a foraging associ- [Traducci6n del autor] auon may exist between the two species.The monkey troopswere alsoattended by the TurkeyVultures. Raptor- ACKNOWLEDGMENTS monkeyassociations usually involve opportunisfic feeding I would like to thank my Venezuelan counterparts, on flushed invertebrates,particularly insects(e.g., Fon- Omar Linares (Universidad Sim6n Bolivar), the lateJos6 taine 1980), and sometimeson vertebratesdisplaced by Luis G6mez Carredano, and more recently Guilberto DECEMBER 2005 SHORT COMMUNICATIONS 461

Rios (Universidad Nacional Experimental de los Llanos FONTAINE,R. 1980. Observationson the foraging associ- Occidentales,Ez•quiel Zamora) for their adviceand sup- ation of Double-toothed Kites and white-faced capu- port; Ivan de Angelisand the late YolandaCarbonell for chin monkeys.Auk 97:94-98. allowing this studyat Hato Las Nieves;Y. Carbonell and GONZALEZBoscAN, V.C. 1987. Los Morichales de los Lla- Alberto Mendoza for their valuable observations; nos Orientales. Ediciones Corpoven, Caracas,Vene- Susanne Renner (Universit/it Ludwig Maximilian, Mu- zuela. nich) for botanical information; and R. Thorstrom, J. Bednarz, and an anonymous referee for their helpful HAVERSCHMIDT,F. 1947. The Black Vulture and Caracara comments on the manuscript. as vegetarians. 49:210. 1962. Notes on the feeding habits and food of LITERATURE CITED some in Surinam. Condor 64:154-158. HOUSTON, D.C. 1988. Competition for food between BALDWIN,J.D. ANDJ.I. BALDWIN.1977. Observationson Neotropicalvultures in forest. Ibis 130:402-417. Cebuscapucinus in southwesternPanama. Primates 18: MCILHENNY,E.A. 1945. An unusual feeding habit of the 937-941. Black Vulture. Auk 62:136-137. BARLOW,C.R. 2004. The utilization of oil-palm kernel by PINTO, O. 1965. Dos frutos da palmeira Elaeisguineenszs Necrosyrtesmonachus in The Gambia. VultureNews 51: na dieta de Cathartesaura ruficollis.Hornero 10:276- 60-62. 277. BORGWO1F PEDERSEN,H. AND H. BALSLEV.1990. Ecuado- ROBINSON,J.G. 1986. Seasonalvariation in use of time rian Palmsfor Agroforestry.AAURep. 23:1-123. and spaceby the wedge-cappedcapuchin monkey, Ce- BRAUN,A. 1997. La Utilidad de las Palmas en Venezuela. busolivaceus: implications for foraging theory. Smith- Fundaci6n Thomas Merle, Carfipano,Venezuela. son. Contrib. Zool. 431:1-60. CRAFTS,R.C., JR. 1968. Turkey Vultures found to feed on ROHL,E. 1949. Fauna descriptivade Venezuela.Bol. Acad coconut. Wilson Bull. 80:327-328. Cienc. Fis. Mat. Nat. 12:1-495. DAWS, D. 1983. Maintenance and social behavior of roost- THIOLLAY,J.-M. 1978. Les rapacesd'une zone de contact ing Turkey Vultures. Pages 322-329 in S.R. Wilbur savane-for6ten C6te d'Ivoire: sp6cialisationsalimen- and J.A. Jackson[EDs.], Vulture biologyand manage- taires. Alauda 46:147-170. ment. University of Press,Berkeley, CA ZHANG,S. AND L. WANG. 2000. Following of brown ca- U.S.A. puchin monkeysby White Hawks in . ELt&S,DJ. AND D. VALENCIA.1982. Unusual feeding be- Condor 102:198-201. haviorby a populationof BlackVultures. Wilson Bull. 94:214. Received19 July 2004; accepted 18 June 2005